Science Reviews 30 (2011) 2309e2328

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Quaternary Science Reviews

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Last straw versus Blitzkrieg overkill: Climate-driven changes in the Arctic Siberian population and the Late extinction problem

P.A. Nikolskiy a,*, L.D. Sulerzhitsky a, V.V. Pitulko b a Geological Institute of the Russian Academy of Sciences, Pyzhevsky per. 7, 119017 Moscow, b Institute for the Material Culture History, Russian Academy of Sciences, Dvortsovaya nab. 18, 191186 Saint Petersburg, Russia article info abstract

Article history: A set of radiocarbon dates on woolly mammoth were obtained from several regions of Arctic Siberia: the Received 8 December 2009 (n ¼ 68), north of the Yana- Lowland (n ¼ 43), and the Taimyr Peninsula Received in revised form (n ¼ 18). Based on these and earlier published dates (n ¼ 201) from the East Arctic, a comparative analysis 22 October 2010 of the time-related density distribution of 14C dates was conducted. It was shown that the frequencies of Accepted 26 October 2010 14C dates under certain conditions reflect temporal fluctuations in mammoth numbers. At the end of the Available online 27 November 2010 Pleistocene the number of in the East Arctic changed in a cyclic manner in keeping with a general “Milankovitch-like” trend. The fluctuations in numbers at the end of the Pleistocene occurred synchronously with paleoenvironmental changes controlled by global climatic change. There were three minima of relative mammoth numbers during the last 50 000 years: 22 000, 14 500e19 000, and 9500 radiocarbon years ago, or around 26 000, 16e20 000, and 10 500 calendar years respectively. The last mammoths lived on the New Siberian Islands, which were connected to the continent at that time, 9470 40 radiocarbon years ago (10 700 70 calendar years BP). This new youngest date approximates the extinction time of mammoths in the last continental refugium of the Holarctic. The adverse combination of environmental parameters was apparently a major factor in the critical reduction in mammoth numbers. The dispersal of humans into the Arctic areas of Siberia no later than 28 000 radiocarbon years ago did not overtly influence animal numbers. Humans were not responsible for the destruction of a sustainable mammoth population. The expanding human population could have become fatal to mammoths during strong the minima of their numbers, one of which occurred at the very beginning of the Holocene. Ó 2010 Elsevier Ltd. All rights reserved.

1. Introduction interspecies and intraspecies competition, disease, and other natural factors. We attribute the astronomical impact hypothesis for The last non-compensated extinctions among large terrestrial Pleistocene extinction to this group of factors, as the impact event mammals started roughly 40 000 years ago, becoming more (if it ever took place) would have operated indirectly, triggering the pronounced towards the end of the Pleistocene. Possible causes of mentioned environmental factors. the extinction have been a topic of discussion for more than 150 2. Extermination by humans. There are many extinction models years. Nearly all of the known hypotheses concerning the disap- in which some of the named factors operate jointly. Different pearance of certain taxa or entire faunas were put forward in the authors have varying ideas regarding the principal model (see for 19th century (Pavlova, 1924; Grayson, 1984). At present, there are example: Martin and Steadman, 1999; Barnosky et al., 2004; two frequently mentioned causes of this extinction (see Barnosky Whitney-Smith, 2004; Wroe et al., 2004; Guthrie, 2006; Wroe et al., 2004; Koch and Barnosky, 2006 for a review): et al., 2006). 1. Transformation of the environment caused by global climatic For a long time, testing these hypotheses has been severely changes, appearing as fluctuations of temperature, humidity, depth hampered by a lack of direct evidence for dependence of animal of snow cover, wind, change of composition and decrease in population sizes upon any particular factor (natural or anthropo- productivity of vegetation, changes in landscapes, soil geochemistry, genic). The recent widespread use of different dating techniques, especially radiocarbon age measurement, has begun to supply the necessary data. * Corresponding author. Tel.: þ7 916 9354872. E-mail addresses: [email protected] (P.A. Nikolskiy), [email protected] Each direct radiocarbon date on a bone, even if not found in situ, (V.V. Pitulko). uniquely corresponds to three parameters:

0277-3791/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.quascirev.2010.10.017 2310 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

1. The animal species (if known). colder climatic conditions would have led to expansion of steppe- 2. The bone location (if known). like landscapes, favorable for abundant and diverse Pleistocene 3. The radiocarbon date of the animal’s death. fauna (Guthrie, 1990; Sher, 1997; Yurtsev, 2001). During interglacial periods, this region served as a refugium for the Pleistocene fauna If the taphonomic biases are low, a sufficient number of dates (Boeskorov, 2006; Lister and Stuart, 2008) because it preserved the allows for reconstruction of the spatio-temporal dynamics of animal conditions of the colder periods of the Pleistocene. Therefore, this ranges and abundance. Furthermore, the radiocarbon method region is one of the world’s largest sources of Late Pleistocene allows for precise dating of some natural and anthropogenic mammal remains. processes. A joint comparative analysis of densely-dated sequences We selected mammoths, Mammuthus primigenius (Blu- is one of the most promising methods to search for cause-effect menbach), as the subject of study because their remains are well relationships of the Late Pleistocene extinction (Martin, 1984; represented in the material collected by the “Zhokhov-Yana” project Guthrie, 1990, 2006; Sulerzhitsky, 1995; Graham et al., 1996; and are the most suitable for conventional radiocarbon dating, Sulerzhitsky and Romanenko 1997; Vasil’chuk et al., 1997; Sher, which requires large samples. In addition, the dates on mammoth 1997; Bocquet-Appel et al., 2005; Orlova et al., 2001; available from the literature, which we included in the analysis, are MacPhee et al., 2002; Stuart et al., 2002; Kuzmin et al., 2003; most abundant. Finally, the mammoth was one of the dominant Barnosky et al., 2004; Gamble et al., 2004; Kuzmin and Orlova, members of the Late Pleistocene (“ Mammoth”) Fauna of the 2004; Orlova et al., 2004; Stuart et al., 2004; Sher et al., 2005; Holarctic and it definitely played a role in the human economy, as Stuart, 2005; Ugan and Bayers, 2007; and many other authors). mammoth remains occur in many Paleolithic sites. The significant By the late 1980s, numerous localities of northeastern Asia number of new reliable dates with accurate sampling, and careful had yielded a large series of 14C dates on mammoth bone collagen analysis of the dating results, enable a study of short-period fluc- (Orlova, 1979; Lavrov and Sulerzhitsky, 1992; Sulerzhitsky, 1995; tuations in frequencies of radiocarbon dates. This makes possible Sulerzhitsky, 1997; Sulerzhitsky and Romanenko, 1997). The anal- a detailed comparison between the dynamics of mammoth ysis of these data revealed a distribution of dates over time resem- numbers, environmental fluctuations and human impact. bling in shape the well-known climatic trend of the terminal Contrary to other regions of Eurasia where the “Mammoth Pleistocene. Most dates are concentrated within the 45e24 ka Fauna” coexisted with people for a longer time, the northern region interval (all dates quoted uncalibrated), which preceded the of the Eastern Arctic was not populated by humans until a relatively beginning of the (LGM). Within the LGM late period. This provides an important opportunity to evaluate the (23e15 ka), the number of dates strongly decreases, followed by “Blitzkrieg overkill” hypothesis for northernmost Asia. The Blitz- a short-term increase. In the interval corresponding to the early krieg overkill hypothesis assumes that people destroyed a sustain- Holocene, the number of dates sharply decreases to zero, indicating able megamammal population shortly after first contact. The the time of mammoth extinction (in Wrangel [Vartanyan et al.,1993, alternative “Last straw” hypothesis assumes a low influence of 1995, 2008; Vartanyan, 2007] and Pribilof Islands (Guthrie, 2004; humans on the sustainable mammal population, but a human- Veltre et al., 2008) woolly mammoths survived somewhat longer). induced extinction of megamammal populations stressed by The subsequent considerable increase in the number of radio- adverse environmental conditions. carbon dates suggests that the general trend remains unchanged (compare Sulerzhitsky, 1997; Sher et al., 2005); and general trends 2. Methods and material (but not details) of date distribution from different areas of Arctic Siberia and northern North America are much alike (compare There are two hypotheses requiring evaluation prior to the Matheus et al. (2003), Guthrie (2004), and Sher et al. (2005)). It was analysis of human vs. naturally-induced causation of mammoth suggested that this stability in the temporal distribution of dates extinction in Arctic Siberia: and its spatial consistency is not coincidental and is indicative of fluctuations in actual numbers at the end of the Pleistocene. At the 1. The distribution of date numbers through time represents same time, it was noticed that major fluctuations in the frequency actual trends in the mammoth population, and is not signifi- of dates broadly correspond to the paleoclimatic pattern and, cantly affected by taphonomic biases. hence, the paleoclimatic changes may be their cause. According to 2. Mammoth numbers depend on climate-driven environmental this approach, the number of dates is a priori regarded as a function conditions (although this statement is broadly accepted, it of animal numbers. Although a few authors (Sulerzhitsky, 1997; remains insufficiently demonstrated). Matheus et al., 2003; Sher et al., 2005; Surovell et al., 2009)have paid special attention to the possible influence of taphonomic, To test the first assumption, we have compared the density geological, and other postmortal factors on date frequency, it is distribution of mammoth-based 14C dates from five regions of clear that dependence of date numbers on animal numbers, the Arctic Siberia remote from each other (Fig. 1). Any marked climate, and taphonomy are relevant to the extinction problem and agreement of these patterns would indicate an effect of some are require special evaluation. We have had the opportunity to uniform parameter, such as the abundance of mammoths and/or conduct such research on abundant material obtained in the New taphonomy. To reveal actual changes in mammoth populations it is Siberian Islands and the adjacent lowland, through the framework very important to estimate the influence of the taphonomy on the of the Russian-American project “Zhokhov-Yana”. distribution of the dates. Taphonomic biases could be introduced by The northern portion of Eastern Siberia has proved to be an ideal large-scale climate-driven factors, such as regional or global cycles region to recover high-quality continuous series of radiocarbon in deposition and exposure of fossil-bearing layers. These could be dates on mammalian remains. At the end of the Pleistocene, this similar across the whole region. In this case taphonomic biases area did not experience the destructive action of continental could contribute significantly to date distribution trends. glaciation. Permafrost conditions, persisting here for a million The evidence suggests a low effect of taphonomy on the years, have allowed the preservation of buried bones and even soft frequency of dates (at least for Beringia): tissues of animals. During the colder periods of the Pleistocene, this area was a part of the extensive Beringian land that incorporated 1. Individual frequency distributions of dates are not equal for vast emerging shelf areas due to sea level falls. At the same time, different mammal taxa (mammoth, bison, horse, caribou, P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2311

Fig. 1. Geographic locations of the areas and individual sites that yielded 14C-dated mammoths discussed in the text. Points designate sampling sites. NS and TM are areas that produced 14C date samples of the “areal” type. OY, B, L, are individual sites yielding series of 14C dates of the “spot” type. NS, New Siberian islands (including , Fad- deevsky, and Kotelny Islands); TM, Taimyr region (including Taimyr Peninsula, lower reaches of the Khatanga River area, and Severnaya Zemlya Archipelago); OY, Oyogos-Yar; B, Bykovsky Peninsula; L, southern coast of Bolshoi Lyakhovsky Island.

muskox) sampled in the New Siberian Islands (Nikolskiy et al., distributions, it is quite probable that the influence of in preparation), Taimyr Peninsula (MacPhee et al., 2002), and taphonomy is low.1 Northern Alaska (Matheus et al., 2003). If taphonomy was 3. One of the largest 14C samples discussed in this report was a significant effect, the frequency distributions of dates of collected at the Mamontova Hayata site, Bykovsky peninsular different taxa would be similar. (Schirrmeister et al., 2002; Sher et al., 2005)(Fig. 1B). This is 2. 14C frequency distributions, and mtDNA data, suggest similar trends in the demographic history of Beringian bison (Shapiro 1 mtDNA study of mammoths suggests no demographic changes before and et al., 2004). Since we can detect the population changes during the LGM. But only 25 samples were analyzed, probably insufficient to 14 (independently confirmed by mtDNA) through C frequency demonstrate trends (Barnes et al., 2007). 2312 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

a very well-documented section of Late Pleistocene deposits. The precise ages of units in the 50-m high section were defined by about one hundred radiocarbon determinations. The distribution of dates over the section suggests roughly equal rates of deposition (Schirrmeister et al., 2002, p. 101). But mammoth 14C date frequencies are not equally distributed (Fig 5B). This most probably suggests some other, not taphonomic biases on mammoth bone distribution.2

Although we believe that taphonomic biases did not contribute significantly to date distribution trends, we applied strict quality- control criteria to the material to minimize bias in the distribution of dates not reflecting fluctuations of animal numbers, but related to features of taphonomy, sedimentation, postsedimentary concentration of bones, and overrepresentation in the sample of bones of a certain age. These criteria include: Fig. 2. Map showing localities where Quaternary mammal remains have been e Material that is sufficiently abundant (hundreds of specimens) collected over the New Siberia Island. This island is one of the biggest among the New to produce a representative, unbiased series of samples for 14C Siberian Islands, and most of the bones for dating have been collected here. Bones are uniformly distributed across the area, do not form marked concentrations, and are not dating (many tens of samples). Smaller collections favor inci- specifically confined to certain local geological bodies. Linear and point concentrations dental enrichment or depletion of certain age intervals. are controlled only by the density of field routes. e Samples have to originate as far as possible from areas with similar environmental conditions. Marked differences in local conditions can influence the number of mammoths, causing combining samples from too-vast areas may result in the loss of a shift in date distribution. weak signals. 14 e Bone remains for dating should be collected from extensive We have selected 330 C dates on mammoth bone collagen. areas in order to balance out the influence of local taphonomic Of these, 192 dates have been obtained directly by us in the Labo- and other selective factors. ratory of Geochemistry of Isotopes and Geochronology of the e Samples should not be overloaded with materials from Geological Institute of the Russian Academy of Sciences by the uniquely rich localities (sections). Even if such a site (section) scintillation (conventional) method (87 dates reported here for fi 3 has sufficient stratigraphic range, it is unlikely to yield an even the rst time). One-hundred and thirty eight additional dates are distribution of dated material throughout this range. taken from the literature. The list of dates with their basic e Samples should not include specimens from archeological sites descriptions is given in Table 1. 14 and natural mass accumulations of animal remains. It is Two sets of C dates of mammoth bones represent areal obvious that both cases will produce artificial peaks within the samples. They closely met the quality criteria listed above and were fi distribution curves corresponding to the age of an archeo- used for the evaluation of the rst hypothesis (the frequency of logical site or the formation time of the mass accumulation. radiocarbon dates of mammoth corresponds to the time-related fi e The collecting should favor a uniform sampling of materials of abundance of this species). The rst set of dates comes from the different age. In general, this is possible when most bones are New Siberian Islands, including New Siberia, Faddeevsky, and sufficiently exposed from . Kotelny Islands (Fig. 1, NS). The second set comes from the Taimyr e Duplication of dates (dating of bones from the same skeleton) region, which comprises the Taimyr peninsula, the lower reaches should be excluded. of the Khatanga River, and the Severnaya Zemlya Archipelago (Fig. 1, TM). fi The first hypothesis (the distribution of date numbers through The NS set includes 86 nite dates from many tens of small 2 time represents actual trends in the mammoth population, and is localities scattered throughout the vast, 35 000 km ,area(Fig. 2). not significantly affected by taphonomic biases) would be addi- The majority of bones were collected on gentle slopes of small tionally indirectly confirmed by dissimilarity between (a) series of rivers and creeks shaped by thermoerosion, in creek channels, and dates that conform to the above criteria, and (b) those that do not, on the coast. fi i.e. that are biased by the factors not related to animal numbers. The TM sample consists of 125 nite dates on mammoth 2 Therefore, we compared dates based on two different types of remains collected from an area covering 250 000 km (Severnaya samples: “areal”, and “spot”. Areal samples (Fig. 1 NS, TM) comprise Zemlia is not included in this area count). The sampling protocols mammoth bones collected over a vast area, assuming that small were the same as in the NS sample (personal observations of effects of local taphonomy would be reduced by mutual compen- Sulerzhitsky; see also MacPhee et al., 2002). sation. Spot samples (Fig 2 B, L, OY) consist of material collected The settings of bone occurrence in both collection areas are over a limited space, usually along an individual outcrops up to similar to each other and differ from conditions in many other several kilometers long. The influence of taphonomy would here be Arctic Siberian regions. These distinctive features include the more apparent, as unequal bone accumulation and subsequent scattered areal occurrence of bones, the lack of major bone fi exposure would be more probable at a local site than across the concentrations, the absence of an evident connection with speci c sum of many sites spread over a large area. At the same time we stratigraphic or lithogical settings, and a nearly homogeneous should not mix material from several areal samples, because in distribution of bones throughout the sequence of Late Pleistocene different regions the population dynamics could vary due to Ice Complexes if found in situ (Fig. 2). This type of occurrence and different environments. It is important to take into account that distribution of bones implies limited or no transportation of the

2 We strongly suspect that postmortem elimination of bones from the record, if 3 The method used for the treatment of bone collagen was established by any, was very low, due to the conserving action of permafrost. Arslanov and Svezhentsev (1993) and modified by Sulerzhitsky (1997). P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2313

Table 1 Radiocarbon age on collagen from mammoth bones in Arctic Siberia New Siberian Islands (without B. Liakhovsky Island).

No. Lab. number Date 14C yr BP Dated Position on Locality Source and remarks bone/tissue the map 1 GIN-11874 9470 40 Tusk Fig. 1 (NS) New Siberia Island This report 2 GIN-11245 9650 60 Tooth Fig. 1 (NS) New Siberia Island Anisimov et al. (2005) 3 GIN-11255 10 200 100 Tooth Fig. 1 (NS) New Siberia Island This reporta 4 GIN-11271v 10 260 120 Tooth Fig. 1 (NS) New Siberia Island This reporta 5 GIN-11088 10 580 50 Tusk Fig. 1 (NS) New Siberia Island This reporta 6 GIN-11087 10 950 50 Tusk Fig. 1 (NS) New Siberia Island This reporta 7 GIN-11263 11 060 40 Limb bone Fig. 1 (NS) New Siberia Island This reporta 8 GIN-11258 12 250 50 Tusk Fig. 1 (NS) New Siberia Island This reporta 9 LU-2096 12 590 60 Tusk Fig. 1 (NS) Verkulich et al. (1989) 10 GIN-11244 13 050 150 Tooth Fig. 1 (NS) New Siberia Island This reporta 11 GIN-11880 13 230 70 Tusk Fig. 1 (NS) New Siberia Island This report 12 GIN-11875 13 250 60 Tusk Fig. 1 (NS) New Siberia Island This report 13 GIN-8230 13 700 100 Bone Fig. 1 (NS) Kotelniy Island Sulerzhitsky and Romanenko (1997) 14 LU-1671 15 420 110 Tusk Fig. 1 (NS) Kotelniy Island Makeyev et al. (1989) 15 GIN-11256 17 500 300 Tooth Fig. 1 (NS) New Siberia Island This reporta 16 GIN-8229 18 500 120 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 17 GIN-11823 18 850 70 Tusk Fig. 1 (NS) New Siberia Island This report 18 GIN-10645 19 400 90 Tibia Fig. 1 (NS) Faddeevsky Island This reportb 19 LU-1790 19 990 110 Tusk Fig. 1 (NS) Kotelniy Island Makeyev et al. (1989) 20 GIN-11271v 20 150 250 Tooth Fig. 1 (NS) New Siberia Island This reporta 21 GIN-10644 20 340 80 Tibia Fig. 1 (NS) Faddeevsky Island This reportb 22 GIN-11878 20 800 70 Tusk Fig. 1 (NS) New Siberia Island This report 23 GIN-5760 20 900 100 Tusk Fig. 1 (NS) Faddeevsky Island Sulerzhitsky (1995) 24 GIN-11254 23 350 500 Tooth Fig. 1 (NS) New Siberia Island This reporta 25 GIN-11253 23 510 120 Tusk Fig. 1 (NS) New Siberia Island This reporta 26 GIN-8244 23 940 150 Tusk Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 27 GIN-11257b 24 800 300 Tooth Fig. 1 (NS) New Siberia Island This reporta 28 GIN-8227 25 180 150 Tusk Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 29 GIN-8246 25 200 180 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 30 GIN-8532 25 540 170 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 31 GIN-4710b 25 800 200 Tusk Fig. 1 (NS) Faddeevsky Island Stuart (1991) 32 GIN-8224 27 100 300 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 33 GIN-11268 27 300 200 Humerus? Fig. 1 (NS) New Siberia Island This reporta 34 GIN-4710 28 000 200 Tusk Fig. 1 (NS) Faddeevsky Island Stuart (1991) 35 GIN-8225 28 650 350 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 36 GIN-11257a 29 000 1000 Tooth Fig. 1 (NS) New Siberia Island This reporta 37 LU-1791 29 020 190 Tusk Fig. 1 (NS) Kotelniy Island Makeyev et al. (1989) 38 GIN-4711 29 100 1000 Rib Fig. 1 (NS) Faddeevsky Island Stuart (1991) 39 GIN-4330 29 100 400 Radius Fig. 1 (NS) Faddeevsky Island Lavrov and Sulerzhitsky (1992) 40 GIN-11252 29 500 300 Tusk Fig. 1 (NS) New Siberia Island This reporta 41 GIN-8260 29 700 250 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 42 GIN-11897 29 850 600 Tusk Fig. 1 (NS) New Siberia Island This report 43 GIN-8236 29 900 300 Bone Fig. 1 (NS) Lopatka Peninsula Sulerzhitsky and Romanenko (1997) 44 GIN-11251 30 400 600 Tooth Fig. 1 (NS) New Siberia Island This reporta 45 GIN-11791 31 100 350 Scapula Fig. 1 (NS) New Siberia Island This report 46 GIN-11271 31 300 700 Tooth Fig. 1 (NS) New Siberia Island This reporta 47 GIN-8226 31 400 300 Tusk Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 48 GIN-11259 31 600 700 Tooth Fig. 1 (NS) New Siberia Island This reporta 49 GIN-11789 31 900 300 Femur Fig. 1 (NS) New Siberia Island This report 50 GIN-11265 33 000 700 Limb bone Fig. 1 (NS) New Siberia Island This reporta 51 GIN-8233 33 600 500 Bone Fig. 1 (NS) Lopatka Peninsula Sulerzhitsky and Romanenko (1997) 52 GIN-11877 33 950 260 Tusk Fig. 1 (NS) New Siberia Island This report 53 GIN-11270 34 200 700 Scapula Fig. 1 (NS) New Siberia Island This reporta 54 GIN-11899 34 400 400 Tusk Fig. 1 (NS) New Siberia Island This report 55 GIN-8254 34 400 400 Bone Fig. 1 (NS) Kotelniy Island Sulerzhitsky and Romanenko (1997) 56 GIN-8247 34 500 500 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 57 GIN-11247 34 560 400 Tusk Fig. 1 (NS) New Siberia Island This reporta (continued on next page) 2314 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Table 1 (continued )

No. Lab. number Date 14C yr BP Dated Position on Locality Source and remarks bone/tissue the map 58 GIN-8243 35 210 500 Tusk Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 59 GIN-11264 35 400 400 Tusk Fig. 1 (NS) New Siberia Island This reporta 60 GIN-11900 35 500 300 Tusk Fig. 1 (NS) New Siberia Island This report 61 GIN-11876 35 600 400 Tusk Fig. 1 (NS) New Siberia Island This report 62 GIN-11028 35 600 400 Tusk Fig. 1 (NS) New Siberia Island This reporta 63 GIN-11691 35 700 600 Mandible Fig. 1 (NS) New Siberia Island This report 64 GIN-8223 35 800 700 Bone Fig. 1 (NS) Bel’kovskiy Island Sulerzhitsky and Romanenko (1997) 65 GIN-8238 36 000 500 Bone Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 66 GIN-11822 36 100 350 Tusk Fig. 1 (NS) New Siberia Island This report 67 GIN-11248 36 600 500 Tusk Fig. 1 (NS) New Siberia Island This reporta 68 GIN-8243a 36 700 500 Tusk Fig. 1 (NS) Faddeevsky Island Sulerzhitsky and Romanenko (1997) 69 GIN-11879 37 150 700 Tusk Fig. 1 (NS) New Siberia Island This report 70 GIN-112672b 38 250 600 Scapula Fig. 1 (NS) New Siberia Island This reporta 71 GIN-11267zh 38 450 600 Scapula Fig. 1 (NS) New Siberia Island This reporta (the same specimen as GIN-112672b) 72 GIN-11250 38 900 500 Tusk Fig. 1 (NS) New Siberia Island This reporta 73 GIN-11820 40 260 500 Tusk Fig. 1 (NS) New Siberia Island This report 74 GIN-11266 40 300 1000 Tusk Fig. 1 (NS) New Siberia Island This reporta 75 GIN-11246 40 500 600 Tusk Fig. 1 (NS) New Siberia Island This reporta 76 GIN-11693 40 900 1000 Tooth Fig. 1 (NS) New Siberia Island This report 77 GIN-11824 40 900 700 Tusk Fig. 1 (NS) New Siberia Island This report 78 GIN-11029 41 000 600 Tusk Fig. 1 (NS) New Siberia Island This reporta 79 GIN-11788 41 350 600 Tusk Fig. 1 (NS) New Siberia Island This report 80 GIN-11031 44 100 1000 Tusk Fig. 1 (NS) New Siberia Island This reporta 81 GIN-11821 44 400 1100 Tusk Fig. 1 (NS) New Siberia Island This report 82 GIN-11030 44 400 1400 Tusk Fig. 1 (NS) New Siberia Island This reporta 83 GIN-11694 45 000 1000 Tusk Fig. 1 (NS) New Siberia Island This report 84 GIN-11787 45 500 1300 Tusk Fig. 1 (NS) New Siberia Island This report 85 GIN-11817 47 100 1500 Tusk Fig. 1 (NS) New Siberia Island This report 86 GIN-11818 48 600 1500 Tusk Fig. 1 (NS) New Siberia Island This report 87 GIN-11262 >33 500 Femur Fig. 1 (NS) New Siberia Island This reporta 88 GIN-11790 >36 900 Tusk Fig. 1 (NS) New Siberia Island This report 89 GIN-11692 >40 000 Limb bone Fig. 1 (NS) New Siberia Island This report 90 GIN-11261 >45 000 Limb bone Fig. 1 (NS) New Siberia Island This reporta 91 GIN-11695 >47 000 Rib and humerus Fig. 1 (NS) New Siberia Island This report 92 GIN-11271a >47 900 Tusk Fig. 1 (NS) New Siberia Island This reporta 93 GIN-11086 >48 000 Tusk Fig. 1 (NS) New Siberia Island This reporta 94 GIN-11260 >48 000 Limb bone Fig. 1 (NS) New Siberia Island This reporta 95 GIN-11819 >50 000 Tusk Fig. 1 (NS) New Siberia Island This report

Oyogos-Yar 1 GIN-11898 16 200 400 Limb bone Fig. 1 (OY) Svyatoy Nos Cape This reporte 2 GIN-9556 17 100 300 Tooth Fig. 1 (OY) Kondrat’eva This reportc 3 GIN-10651 19 800 130 Radius Fig. 1 (OY) Svyatoy Nos Cape This reporte 4 GIN-10560 22 700 700 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 5 GIN-10561 23 000 600 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 6 GIN-9045 25 200 300 Limb bone Fig. 1 (OY) Svyatoy Nos Cape This reportd 7 GIN-9563 26 100 600 Tusk Fig. 1 (OY) Kondrat’eva This reportc 8 GIN-10594 27 000 600 Tooth Fig. 1 (OY) Svyatoy Nos Cape This reporte 9 GIN-10600 29 100 300 Tooth Fig. 1 (OY) Svyatoy Nos Cape This reporte 10 GIN-10652 34 400 400 Ulna Fig. 1 (OY) Svyatoy Nos Cape This reporte 11 GIN-10564 34 500 800 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 12 GIN-9558 36 400 400 Tusk Fig. 1 (OY) Kondrat’eva This report 13 GIN-9565 37 200 400 Tusk Fig. 1 (OY) Kondrat’eva This reporte 14 GIN-10657 40 500 900 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 15 GIN-10555 40 800 1500 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 16 GIN-9566 40 900 1200 Tusk Fig. 1 (OY) Ulakhan-Taala This reportc 17 GIN-9552 42 900 900 Limb bone Fig. 1 (OY) Ulakhan-Taala This reportc 18 GIN-9568 43 100 1000 Tusk Fig. 1 (OY) Kondrat’eva This reportc 19 GIN-9557 47 400 1200 Tusk Fig. 1 (OY) Kondrat’eva This reportc 20 GIN-9044 48 800 140 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reportd 21 GIN-9555 >16 000 Limb bone Fig. 1 (OY) Kondrat’eva This reportc 22 GIN-10655 >16 700 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 23 GIN-9564 >31 600 Tusk Fig. 1 (OY) Ulakhan-Taala This reporte 24 GIN-10596 >34 700 Tooth Fig. 1 (OY) Svyatoy Nos Cape This reporte 25 GIN-10602 >34 700 Tooth Fig. 1 (OY) Svyatoy Nos Cape This reporte 26 GIN-9559 >36 900 Tusk Fig. 1 (OY) Kondrat’eva This reportc 27 GIN-10563 >38 200 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 28 GIN-10601 >38 600 Tooth Fig. 1 (OY) Svyatoy Nos Cape This reporte 29 GIN-11901 >40 900 Tooth Fig. 1 (OY) Svyatoy Nos Cape This reporte P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2315

Table 1 (continued )

No. Lab. number Date 14C yr BP Dated Position on Locality Source and remarks bone/tissue the map 30 GIN-10559 >44 200 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 31 GIN-10565 >44 200 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 32 GIN-10557 >44 900 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 33 GIN-9567 >46 000 Tusk Fig. 1 (OY) Kondrat’eva This reporte 34 GIN-10556 >46 400 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 35 GIN-10597 >48 400 Tusk Fig. 1 (OY) Aebeliakh Bay This reporte 36 GIN-10598 >48 900 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 37 GIN-9560 >49 000 Tusk Fig. 1 (OY) Ulakhan-Taala This reporte 38 GIN-9569 >49 000 Tusk Fig. 1 (OY) Kondrat’eva This reporte 39 GIN-9562 >50 000 Tusk Fig. 1 (OY) Kondrat’eva This reporte 40 GIN-10653 >50 000 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 41 GIN-10558 >50 200 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 42 GIN-10595 >50 200 Tusk Fig. 1 (OY) Svyatoy Nos Cape This reporte 43 GIN-9561 >51 000 Tusk Fig. 1 (OY) Ulakhan-Taala This reporte

Taimyr Peninsula region 1 GIN-1828 9670 60 Tusk Fig. 1 (TM) Taimyr e Nizhnyaya Stuart (1991) Taimyra R 2 GIN-8256 9780 40 Tusk Fig. 1 (TM) Taimyr, north-east coast Sulerzhitsky and Romanenko (1997) 3 GIN-1495 9860 50 Tooth Fig. 1 (TM) Taimyr e Nizhnyaya Taimyra R Lavrov and Sulerzhitsky (1992) 4 GrA-17350 9920 60 Mandible Fig. 1 (TM) Taimyr e Bikada R MacPhee et al. (2002) 5 GIN-10508 10 070 60 Tusk Fig. 1 (TM) Taimyr e Nyen’gatiatari R MacPhee et al. (2002) 6 GIN-1489 10 100 100 Tooth Fig. 1 (TM) Taimyr e Engelhardt L Lavrov and Sulerzhitsky (1992) 7 GIN-11138a 10 200 40 Tooth Fig. 1 (TM) Taimyr e Kupchiktakh L MacPhee et al. (2002) 8 GIN-10507 10 270 120 Tibia Fig. 1 (TM) Taimyr e Nyun’karakutari R MacPhee et al. (2002) 9 Beta-148640 10 270 40 Tusk Fig. 1 (TM) Taimyr e Gol’tsovaya R MacPhee et al. (2002) 10 GIN-1828k 10 300 100 Radius Fig. 1 (TM) Taimyr e Nizhnyaya Taimyra R Stuart (1991) 11 LuA-4820 10 450 110 Tusk Fig. 1 (TM) Severnaya Z, Möller et al. (2006) October Revolution Is. 12 GIN-3768 10 680 70 Limb bone Fig. 1 (TM) Taimyr e Taimyr L, Stuart (1991) Nganasanskaya R 13 GIN-10552 10 790 100 Tooth Fig. 1 (TM) Taimyr e Krasnaya R MacPhee et al. (2002) 14 GIN-11529 10 950 70 Atlas Fig. 1 (TM) Taimyr This reportg 15 GIN-3067 11 140 180 Tooth Fig. 1 (TM) Taimyr e Taimyr L, Baikura Neru B Stuart (1991) 16 T-297 11 450 250 Soft tissues Fig. 1 (TM) Taimyr e Mamonta R Heintz and Garutt (1964) 17 LU-610 11 500 60 Tusk Fig. 1 (TM) Severnaya Z e Makeev et al. (1979) October Revolution Is. 18 GIN-8881 11 870 180 Bone Fig. 1 (TM) Taimyr e Taimyr L, Cape Sabler Dereviagin et al. (1999) 19 Beta-148663 11 940 40 Ulna Fig. 1 (TM) Taimyr e Arylakh L MacPhee et al. (2002) 20 GIN-10506 12 050 100 Pelvis? Fig. 1 (TM) Taimyr This reportf 21 GIN-10506 12 050 100 Tibia Fig. 1 (TM) Taimyr e Bikada R MacPhee et al. (2002) 22 GIN-1783 12 100 80 Limb bone Fig. 1 (TM) Taimyr e Taimyr L, Stuart (1991) Baskura Peninsula 23 GIN-2943r 12 260 120 Limb bone Fig. 1 (TM) Taimyr e Severnaya R Lavrov and Sulerzhitsky (1992) 24 GIN-3242 12 450 120 Limb bone Fig. 1 (TM) Taimyr e Severnaya R Lavrov and Sulerzhitsky (1992) 25 GIN-2677 12 780 80 Limb bone Fig. 1 (TM) Taimyr e Bikada R Stuart (1991) 26 LU-3827 13 200 130 Tusk Fig. 1 (TM) Taimyr e Taimyr L, Möller et al. (1999) Cape Sabler 27 GIN-2758a 13 340 240 Limb bone Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 28 Beta-148636 13 560 40 Fig. 1 (TM) Khatanga R. basin MacPhee et al. (2002) 29 GIN-8883 13 620 70 Bone Fig. 1 (TM) Taimyr e Taimyr L. Dereviagin et al. (1999) 30 GIN-3518 14 800 50 Bone Fig. 1 (TM) Khatanga e Khatanga R basin, Sulerzhitsky et al. (1986) Ulakhan-Yuryakh R 31 Beta-148642 15 390 50 Femur Fig. 1 (TM) Taimyr e Taimyr L, MacPhee et al. (2002) Cape Sabler 32 GIN-3130 16 330 100 Mandibula Fig. 1 (TM) Taimyr e Stuart (1991) Bolshaya Balakhnya R 33 Beta-148646 18 190 60 Molar Fig. 1 (TM) Taimyr e MacPhee et al. (2002) Bolshaya Balakhnya R 34 LU-654A 19 640 330 Tusk Fig. 1 (TM) Severnaya Z e Makeev et al. (1979) October Revolution Is. 35 LU-688 19 970 110 Tooth Fig. 1 (TM) Severnaya Z e Makeev et al. (1979) October Revolution Is., Ushakova R 36 UtC-8138 20 380 140 Hair Fig. 1 (TM) Taimyr e Buigues and Mol (1999) Bolshaya Balachnya River 37 GIN-3952 20 400 100 Tusk Fig. 1 (TM) Taimyr e Dudypta R Sulerzhitsky et al. (1987) 38 Beta-148647 20 620 70 Limb bone Fig. 1 (TM) Taimyr e Mol et al. (2001a, b) Verkhnaya Taimyra River 39 Beta-148633 20 800 70 Mandible Fig. 1 (TM) Taimyr MacPhee et al. (2002) 40 GrA-17604 20 950 190 Calcaneus Fig. 1 (TM) Taimyr e Taimyr L, MacPhee et al. (2002) Baskura Peninsula (continued on next page) 2316 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Table 1 (continued )

No. Lab. number Date 14C yr BP Dated Position on Locality Source and remarks bone/tissue the map 41 GIN-5574 22 000 200 Tooth Fig. 1 (TM) Khatanga Bay area e Stuart (1991) Popigay R basin 42 GIN-3089 22 750 150 Bone Fig. 1 (TM) Taimyr e Taimyr L, Sulerzhitsky (1995) Baskura Peninsula 43 GIN-2763a 23 500 300 Tusk Fig. 1 (TM) Taimyr e Bederbo-Tarida R Stuart (1991) 44 GIN-11530 23 600 300 Tusk Fig. 1 (TM) Taimyr This reportb 45 GIN-1296B 23 800 400 Limb bone Fig. 1 (TM) Taimyr e Taimyr L, Sabler Cape Stuart (1991) 46 Beta-148639 24 170 110 Skull Fig. 1 (TM) Taimyr e Trautfetter R MacPhee et al. (2002) 47 Beta-148651 24 250 110 Skull Fig. 1 (TM) Khatanga e Munchirdakh L MacPhee et al. (2002) 48 GIN-8737 24 760 110 Scapula Fig. 1 (TM) Taimyr e Labaz L Sher et al. (2005) 49 GIN-2160 24 900 500 Pelvis Fig. 1 (TM) Taimyr e Taimyr L, Baskura Peninsula Lavrov and Sulerzhitsky (1992) 50 GIN-8884 24 990 260 Bone Fig. 1 (TM) Taimyr e Taimyr L, Sabler Cape Dereviagin et al. (1999) 51 LU-749B 25 030 210 Tibia Fig. 1 (TM) Severnaya Z e October Revolution Is., Makeev et al. (1979) Knizhnaya R 52 LE-612 25 100 550 Soft tissues Fig. 1 (TM) Taimyr e Pyasina R Zubakov and Kind (1974) 53 Beta-148634 25 800 130 Fig. 1 (TM) Khatanga R. basin MacPhee et al. (2002) 54 Beta-148665 26 100 170 Humerus Fig. 1 (TM) Taimyr e Taimyr L, Baskura Peninsula MacPhee et al. (2002) 55 GIN-11127 26 200 150 Tusk Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R MacPhee et al. (2002) 56 GIN-1216 26 700 700 Tusk Fig. 1 (TM) Khatanga e Putorana Plt., Stuart (1991) Maymecha R basin 57 GIN-11510 26 900 250 Humerus Fig. 1 (TM) Taimyr This reportf 58 GIN-3836 27 300 200 Rib Fig. 1 (TM) Taimyr e Logata R Stuart (1991) 59 GIN-3929 27 500 300 Tusk Fig. 1 (TM) Taimyr e Kubalakh R Sulerzhitsky et al. (1987) 60 Beta-210777 27 740 220 Mandible Fig. 1 (TM) Taimyr e Baikura-Turku Bay, Poinar et al. (2006) Lake Taimyr 61 Beta-148662 28 270 210 Ulna Fig. 1 (TM) Taimyr MacPhee et al. (2002) 62 Beta-148643 28 310 170 Molar Fig. 1 (TM) Taimyr e Taimyr L, coast opposite MacPhee et al. (2002) Kupffer Is. 63 GIN-952 28 800 600 Tusk Fig. 1 (TM) Taimyr e Shrenk R Kind et al. (1978) 64 GIN-5073 28 900 300 Tusk Fig. 1 (TM) Khatanga R. basin e Anabarka R Lavrov and Sulerzhitsky (1992) 65 GIN-8882 29 250 450 Bone Fig. 1 (TM) Taimyr e Taimyr L, Sabler Cape Dereviagin et al. (1999) 66 GIN-2155 29 500 300 Tusk Fig. 1 (TM) Taimyr e Taimyr L, Matuda P Stuart (1991) 67 GIN-2154 29 500 350 Tooth Fig. 1 (TM) Taimyr This report 68 Beta-148635 29 990 280 Humerus Fig. 1 (TM) Khatanga e Talalakh L MacPhee et al. (2002) 69 GIN-11130a 30 850 200 Mandible Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R MacPhee et al. (2002) 70 Beta-148637 30 890 290 Tooth Fig. 1 (TM) Khatanga e Talalakh L MacPhee et al. (2002) 71 GIN-11504 31 300 200 Tusk Fig. 1 (TM) Taimyr This reportf 72 Beta-148631 31 580 330 Tibia Fig. 1 (TM) Taimyr e Baikuraturku L MacPhee et al. (2002) 73 GIN-3240a 31 800 500 Skull Fig. 1 (TM) Taimyr e Severnaya River 74 GIN-3117 32 000 200 Femur Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Stuart (1991) 75 GIN-2151 32 000 500 Limb bone Fig. 1 (TM) Taimyr e Taimyr L, Matuda P Stuart (1991) 76 GIN-11137 32 200 800 Tooth Fig. 1 (TM) Taimyr e Kupchiktakh L MacPhee et al. (2002) 77 GIN-5074 32 300 400 Tusk Fig. 1 (TM) Khatanga basin area e Popigay R Lavrov and Sulerzhitsky (1992) 78 Beta-148667 32 530 270 Skull Fig. 1 (TM) Taimyr e Taimyr L, Sabler Cape MacPhee et al. (2002) 79 GIN-8261 32 600 700 Bone Fig. 1 (TM) Taimyr e Pyasina R, Farvaterniy Is Sulerzhitsky and Romanenko (1997) 80 Beta-148632 32 750 280 Tibia Fig. 1 (TM) Taimyr MacPhee et al. (2002) 81 Beta-148641 32 840 290 Femur Fig. 1 (TM) Taimyr e Arylakh L MacPhee et al. (2002) 82 GIN-3821 35 000 500 Femur w/ Fig. 1 (TM) Taimyr e Logata R Lavrov and Sulerzhitsky (1992) marrow 83 GIN-3822 36 200 500 Tusk Fig. 1 (TM) Taimyr e Logata R Lavrov and Sulerzhitsky (1992) 84 GIN-5751 36 600 500 Fig. 1 (TM) Khatanga R basin, e Anabarka R Lavrov and Sulerzhitsky (1992) 85 GIN-3122 36 800 500 Tusk Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 86 Beta-148630 36 950 450 Mandible Fig. 1 (TM) Taimyr e Arylakh L MacPhee et al. (2002) 87 GIN-11137a 37 000 400 Tusk Fig. 1 (TM) Taimyr This reportf 88 GIN-5750 37 000 500 Tooth Fig. 1 (TM) Khatanga e Semieriskyay R Lavrov and Sulerzhitsky (1992) 89 Beta-148666 37 080 460 Fig. 1 (TM) Khatanga e Popygai R MacPhee et al. (2002) 90 GIN-11533 37 200 400 Limb bone Fig. 1 (TM) Taimyr This reportg 91 GIN-942 38 000 1500 Tusk Fig. 1 (TM) Khatanga R, Krestovskiy Mys Kind et al. (1978) 92 GIN-3817 38 300 600 Bone Fig. 1 (TM) Taimyr e Logata R Sulerzhitsky (1995) 93 GIN-11513 38 300 700 Humerus Fig. 1 (TM) Taimyr This reportf 94 GIN-3118 38 400 700 Radius Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 95 GIN-27636 38 500 500 Tooth Fig. 1 (TM) Taimyr e Bederbo-Tarida R Lavrov and Sulerzhitsky (1992) 96 GIN-3136 38 500 600 Femur Fig. 1 (TM) Taimyr e Bederbo-Tarida R Lavrov and Sulerzhitsky (1992) 97 GIN-1491 38 800 1300 Tusk Fig. 1 (TM) Taimyr e Trautfetter R Lavrov and Sulerzhitsky (1992) 98 GIN-3476 38 800 400 Tusk Fig. 1 (TM) Taimyr e Nemu-Dika-Tarida River Sulerzhitsky et al. (1986) 99 GIN-3831 38 900 600 Tusk Fig. 1 (TM) Taimyr e Logata R Lavrov and Sulerzhitsky (1992) 100 Beta-148664 39 050 580 Pelvis Fig. 1 (TM) Taimyr e Arylakh L MacPhee et al. (2002); 101 GIN-3120/II 39 100 1000 Ulna Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 102 GIN-3121/II 39 200 700 Limb bone Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 103 GIN-3071 39 300 500 Femur w/ Fig. 1 (TM) Taimyr e Taimyr L, Baikura Neru B Lavrov and Sulerzhitsky (1992) marrow 104 GIN-11127a 39 300 600 Tusk Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R MacPhee et al. (2002) 105 Beta-148638 39 560 910 Mandible Fig. 1 (TM) Taimyr e Trautfetter R MacPhee et al. (2002) 106 GIN-3135 39 800 600 Skull Fig. 1 (TM) Taimyr e Bederbo-Tarida R Lavrov and Sulerzhitsky (1992) P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2317

Table 1 (continued )

No. Lab. number Date 14C yr BP Dated Position on Locality Source and remarks bone/tissue the map 107 GIN-11134 40 200 600 Sacrum Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R MacPhee et al. (2002) 108 GIN-3804 40 200 600 Tusk Fig. 1 (TM) Taimyr e Logata R Lavrov and Sulerzhitsky (1992) 109 GIN-5025 40 300 400 Tooth Fig. 1 (TM) Khatanga R. basin e Anabarka R Lavrov and Sulerzhitsky (1992) 110 GIN-12710 40 300 600 Limb bone Fig. 1 (TM) Taimyr This reportf 111 GIN-1818/II 40 500 800 Limb bone Fig. 1 (TM) Taimyr e Engelgardt L Lavrov and Sulerzhitsky (1992) 112 Beta-148645 40 560 700 Pelvis Fig. 1 (TM) Taimyr e Taimyr L, Sabler Cape MacPhee et al. (2002) 113 GIN-3100 40 600 600 Limb bone Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 114 GIN-2744B 41 200 1000 Limb bone Fig. 1 (TM) Taimyr e Bederbo-Tarida R Lavrov and Sulerzhitsky (1992) 115 GIN-5224 41 900 800 Tusk Fig. 1 (TM) Khatanga R. basin e Anabarka R Lavrov and Sulerzhitsky (1992) 116 Beta-148648 42 420 500 Tibia Fig. 1 (TM) Taimyr e Arylakh L. MacPhee et al. (2002) 117 GIN-3946 42 800 800 Bone Fig. 1 (TM) Khatanga R. basin e Massonov R Sulerzhitsky et al. (1987) 118 GIN-3072 43 500 1000 Humerus Fig. 1 (TM) Taimyr e Taimyr L, Lavrov and Sulerzhitsky (1992) Baikura Neru B 119 GIN-766 45 000 1000 Limb bone Fig. 1 (TM) Taimyr e Kheta R Lavrov and Sulerzhitsky (1992) 120 GIN-3073 46 100 1200 Scapula Fig. 1 (TM) Taimyr e Taimyr L, Lavrov and Sulerzhitsky (1992) Baikura Neru B 121 GIN-11512 46 400 1500 Limb bone Fig. 1 (TM) Taimyr This reportf 122 GIN-11514 47 200 1400 Tusk Fig. 1 (TM) Taimyr This reportf 123 Beta-148669 47 660 1650 Tooth Fig. 1 (TM) Khatanga e Popygai R MacPhee et al. (2002) 124 GIN-3118a 47 900 1600 Tusk Fig. 1 (TM) Taimyr e Bolshaya Balakhnya R Lavrov and Sulerzhitsky (1992) 125 GIN-689 49 700 1100 Tusk Fig. 1 (TM) Khatanga e Putorana Plt., Lavrov and Sulerzhitsky (1992) Maymecha R 126 GIN-11508 >38 500 Limb bone Fig. 1 (TM) Taimyr This reportf 127 GIN-11128 >38 600 Rib Fig. 1 (TM) Taimyr This reportf 128 GIN-11502 >45 000 Limb bone Fig. 1 (TM) Taimyr This reportf 129 GIN-11509 >45 000 Humerus Fig. 1 (TM) Taimyr This reportf 130 GIN-11505 >49 100 Tusk Fig. 1 (TM) Taimyr This reportf 131 GIN-12160 >49 300 Limb bone Fig. 1 (TM) Taimyr This reportf

Bolshoy Lyakhovsky Island 1 GIN-10713 12 030 60 Ulna Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 2 GIN-10716 12 500 50 Tusk Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 3 GIN-11084 20 800 100 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Andreev et al. (2004) southern coast 4 GIN-10707 22 100 1000 Pelvis Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 5 GIN-10708 25 900 600 Pelvis Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 6 GIN-10706 28 000 180 Ulna Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 7 MAG-316 32 100 900 Skin Fig. 1 (L) Bolshoy Lyakhovsky Island, Lozhkin (1977) southern coast 8 GIN-10659 32 500 500 Vertebrae Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 9 GIN-8739 32 500 1000 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 10 GIN-11085 33 000 320 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Andreev et al. (2004) southern coast 11 GIN-8740 34 800 1000 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003) southern coast 12 GIN-8741 35 100 1800 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003) southern coast 13 GIN-8738 36 500 1600 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003) southern coast 14 GIN-10660 37 800 900 Vertebrae Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 15 GIN-11705 38 700 500 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Andreev et al. (2004) southern coast 16 GIN-10714 39 600 1000 Femur Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 17 GIN-10703 40 200 900 Humerus Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 18 GIN-8813 42 700 1300 Bone Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 19 GIN-10717 43 600 1000 Scapula Fig. 1 (L) Bolshoy Lyakhovsky Island, Kuznetsova et al. (2003), southern coast Sher et al. (2005) 20 OxA-17111 50 200 900 Hair Fig. 1 (L) Bolshoy Lyakhovsky Island, Gilbert et al. (2007) southern coast

Bykovsky Peninsula 1 GIN-10267 12 770 170 Carpale Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 2 GIN-10242 13 100 500 Pelvis Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) (continued on next page) 2318 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Table 1 (continued )

No. Lab. number Date 14C yr BP Dated Position on Locality Source and remarks bone/tissue the map 3 KIA-10679 14 220 80 Skull Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 4 KIA-10680 14 320 60 Rib Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 5 GIN-9881 14 340 280 Metacarpale Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 6 GIN-10239 14 500 110 Scapula Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 7 GIN-10240 14 500 140 Pelvis Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 8 GIN-10234 14 600 100 Rib Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 9 GIN-9907 14 730 60 Pelvis Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 10 GIN-10236 20 200 100 Scapula Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 11 GIN-10248 20 800 600 Metacarpale Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 12 LU-1328 21 630 240 Carpale Fig. 1 (B) Bykovsky Peninsula Tomirdiaro et al. (1984) 13 GIN-9893 23 050 120 Mandible Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 14 GIN-9908 23 100 1200 Metacarpale Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 15 GIN-10264 24 300 200 Vertebra Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 16 GIN-9887 24 800 90 Femur Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 17 GIN-9880 25 000 300 Carpale Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 18 GIN-9882 26 500 230 Femur Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 19 GIN-9896 26 700 900 Pelvis Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 20 GIN-10262 27 400 800 Limb bone Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 21 GIN-10238 28 200 900 Epistropheum Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 22 GIN-9906 28 700 800 Ulna Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 23 GIN-9904 28 800 180 Tusk Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 24 GIN-10247 28 900 200 Pelvis Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 25 GIN-10245 29 400 600 Rib Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 26 GIN-10244 30 300 600 Ulna Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 27 GIN-10260 30 600 900 Ulna Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 28 GIN-10249 31 500 650 Metatarsale Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 29 GIN-9901 32 500 300 Tibia Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 30 GIN-10266 32 800 800 Femur Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 31 GIN-9884 33 000 450 Tibia Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 32 GIN-9890 33 610 300 Tusk Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 33 GIN-10263 33 800 260 Tibia Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 34 GIN-10261 34 000 500 Limb bone Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 35 GIN-9895 34 300 320 Tusk Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 36 GIN-9897 34 900 400 Pelvis Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 37 GIN-9898 35 500 500 Tusk Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 38 GIN-9892 35 700 700 Humerus Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 39 T-171(2) 35 800 1200 Soft tissue Fig. 1 (B) Bykovsky Peninsula Heintz and Garutt (1964) 40 GIN-9900 44 500 1000 Atlas Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002) 41 GIN-9891 44 500 1100 Tibia Fig. 1 (B) Bykovsky Peninsula Schirrmeister et al. (2002)

a Material collected by Anisimov and Tumskoy. b Material collected by Romanenko. c Originally published in Nikolaev et al. (2000). d Originally published in Nikolskiy et al. (1999). e Originally published in Nikolskiy and Basilyan (2003). f Material collected by Koriagin. g Material collected by Orlov. remains from the sites of death. A considerable quantity of bones recording the number of observations within an interval of 10% of was preserved due to permafrost conditions. Active thermoerosion the horizontal axis as it is moved along the x-axis, and weighting in Arctic islands and peninsulas then exposed a significant number them using the cosine function. of bones. These conditions of taphonomy, preservation, and expo- For plotting frequency histograms, samples were empirically sure of bones are optimal for investigating time-related animal divided into classes of 1200 years, which on the one hand would number fluctuations. preserve important peaks of density distributions, and on the other In order to evaluate the influence of local taphonomic and hand minimizes noise. In addition, it is worth noting that the geological conditions on the time-distribution of dates, we have selected histogram interval exceeds the instrumental error of compared the areal samples (NS, TM) with three spot samples. The radiocarbon age measurements which in the oldest dates can latter material came from the following sites: Oyogos-Yar, south amount to thousand years and more. side of Dmitri Laptev Strait e Fig. 1 OY, original data; Mamontova We use Pearson correlation coefficients (r) for statistical Khayata, Bykovsky Peninsula, coast e Fig. 1B(Heintz and comparisons between samples. For each 1200-year bin we take the Garutt, 1964; Tomirdiaro et al., 1984; Schirrmeister et al., 2002); number of dates in one of the comparative samples and the number and Maloe Zimovye, Bolshoi Lyakhovsky Island, northern side of of dates in the other, and these are the paired x, y values. We Dmitri Laptev Strait (Fig. 1 L) (Kuznetsova et al., 2003; Sher et al., examine the probability value (p) to test if the correlation is 2005). significant. For all cases we additionally use visual matching of For the analysis we used both uncalibrated and calibrated dates diagrams for comparisons between samples. when appropriate. To calibrate dates we have applied the Cal- The validity of the first hypothesis (number of dates reflects the Pal_2007 software developed by Weninger et al. (2009). number of animals) makes it possible to evaluate the second We used a density trace and a frequency histogram techniques hypothesis (number of animals is climate-related). For this we have for showing the variation in frequency (concentration) of data compared the observed distributions of radiocarbon dates with points through time. The density trace plots were constructed by a series of high-resolution climatic proxies, as follows: P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2319

2.1. Record of changes in insect faunas from Late Pleistocene record (Weinberg et al., 1999) as a very sensitive, high-resolution deposits of the River delta and continuous climatic proxy.

The recently published record of changes in insect faunas, from 2.2.3. Greenland ice-core data about 50 ka to the beginning of the Holocene, is based on materials Greenland ice-core data have proven to be one of the most from the section of Late Quaternary deposits at the Mamontova important sources of information on past climate. Ice-core isotope Hayata site, Bykovsky Peninsula, the Lena River delta, Arctic Siberia record sensitively responded to temperature fluctuations during (Fig. 1B) (Sher et al., 2005). Fossil entomofaunas are very sensitive the last glacial cycle (Dansgaard et al., 1993; Bender et al., 1994; indicators of environmental changes (vegetation, annual temper- Meese et al., 1994a,b). Strong similarities in oxygen isotope record ature, humidity) in the Pleistocene (Berman and Alfimov, 1993; in six Greenland ice-cores (Camp Century, Dye-3, GRIP, GISP2, Elias, 1999; Kuzmina, 2001; Alfimov et al., 2003; Kuzmina, 2003). Renland and NorthGRIP) have recently been revealed (Johnsen All studied insect species are extant. Data on modern ranges and et al., 2001). This suggests that the dominant influence on oxygen environmental conditions allow us to reconstruct the relative isotope variations reflected in the ice-sheet records was climatic changes of environment during the specified time interval. The change. We have chosen the GISP2 d18O values record (Grootes analysis of insects from the Bykovsky section is aided by a parallel et al., 1993; Meese et al., 1994a,b; Stuiver and Grootes, 2000) with study of palynology and plant remains. A sufficient number of dates age model tuned to Hulu cave data (Weninger and Joöris, 2008). (about 100) and their mutual control help validate the accuracy of the correlation between age and climatic indicators. The record of 2.2.4. Mg/Ca ratio in marine ostracod shells from the North Atlantic fossil insects is sufficiently detailed and contains only a few gaps. Dwyer et al. (2002) reconstructed time-series of last-glacial The Bykovsky Peninsula (Fig. 1) is located just between the New cycle deep-sea temperature from water cores from the Siberian Islands and Taimyr Peninsula, which makes the place ideal western North Atlantic using Mg/Ca ratios of benthic ostracod for studying regional past climatic changes to compare with shells. Mg/Ca analysis suggests that multiple-shell average Mg/Ca mammoth number fluctuations. ratios provide reliable estimates of bottom-water temperature history. The water temperature records show glacial-to-interglacial 2.2. Pollen and insect record in lower Yana River, Arctic Siberia variations, and changes during the deglaciation and the Holocene. The ostracod Mg/Ca ratio study provides evidence that deep-ocean A high-resolution paleoclimatic archive comes from Lower circulation is closely linked to climate change over a range of Yana Paleolithic site at 70430N, 135 250E, Yana River, Arctic timescales regardless of the mean climate state. Thus, benthic Siberia (Pitulko et al., 2004). This well radiocarbon-dated pollen Mg/Ca ratios are a good proxy for high-resolution reconstruction of and insect record spanning 34 000 years was another climatic past climate conditions. proxy we used to compare with mammoth date distributions. Agreement between these very differently originating climatic Based on the environment encompassing the current distribution records and mammoth 14C date frequencies would strongly suggest of plant taxa (the environmental envelope), Pitulko et al. (2007) the dependence of past animal numbers on climate. and Pavlova et al. (2009) reconstructed the average tempera- tures of the warmest month and mean precipitation. Floristic data 3. Results were controlled by paleoentomological proxies. This reconstruc- tion of the paleoenvironmental conditions in terms of absolute 3.1. Comparison between frequency distributions of 14C dates on temperature and moisture is important as it make it possible to mammoths from different regions of Arctic Siberia estimate the response of mammoth population to these climatic factors. 3.1.1. New Siberian Islands versus Taimyr region The frequency distributions of 14C dates of the NS and TM series 2.2.1. Lake El’gygytgyn (Chukotka, Russian Far East) climatic proxy are shown in Fig. 3. Most parts of the distributions are clearly in records close agreement: correlation coefficient is 0.89, p < 0.05 at two El’gygytgyn meteorite impact lake (67300N, 172050E, Russian scale segments 13 000e29 000 and 46 000e48 000 14C years BP, Far East) is the largest and oldest unglaciated lake basin in the which covers 45% of the curve; correlation coefficient is 0.79, Arctic. Its sediments represent a 3.6 million-year-long, mostly time- p < 0.05 if we exclude the segment between 31700 and 41000 ybp continuous record of late Cenozoic Arctic climate evolution. We which reduces the overall correlation because of the weak negative have used for comparison the magnetic susceptibility record during correlation seen in this short fragment. The coefficient correlation the last glacial cycle as a proxy for changes in past temperature of frequencies without any removals is 0.21, p < 0.05. (Asikainen et al., 2007; Brigham-Grette et al., 2007). Magnetic We interpret this correlation as an effect resulting from the susceptibility is a proxy for changes in anoxia driven by changes in same influence, i.e. uniform fluctuation of animal numbers in two seasonal lake ice cover, in turn a function of temperature and extensive regions of Arctic Siberia during the last millennia of the insolation (Asikainen et al., 2007). Pleistocene.

2.2.2. Lake Baikal (South eastern Siberia) climatic proxy record 3.1.2. New Siberian Islands and Taimyr region versus spot-sample- Core data from Lake Baikal form one of the most promising based frequency distributions sources of information on the paleoenvironments of northern Asia. We assume that the comparison of areal and spot samples The continuity of the paleoclimatic record, its sensitivity to orbital should reflect the degree of local taphonomic influences on the forcing, and its high resolution make it a candidate for a “paleo- distribution patterns. All three spot series (B, OY, L) show marked climatic stratotype” section for continental Asia (Prokopenko et al., differences from the areal, NS and TM, series (Fig. 4A, 5B, C). 2000). Climatic cycles are recorded in Lake Baikal as changes of Considering different segments across the six pairwise productivity seen in abundance, organic carbon accumu- comparisons (Fig. 4), the correlation coefficients vary from 0.22 lation, changes in biogenic silica concentrations, sponge spicule to 0.52; moreover, that fraction of the correlation coefficients concentration, lithology, and some geophysical and isotope which are relatively high (0.4e0.5) never reaches more than 20%. parameters. We have chosen the sponge spicule concentration Thus, it is obvious that the local sites (OY, B, and L), even if they 2320 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Fig. 3. Comparison of time-related densities of mammoth-based 14C dates from the New Siberian Islands and the Taimyr region. (A) Fragments of curves with maximal correlation coefficient (r ¼ 0.89, p < 0.05). (B) Fragment of curves where correlation coefficient is lower, but still is very considerable (r ¼ 0.79, p < 0.05). (C) The total curve length correlated: correlation coefficient is weak (r ¼ 0.21, p < 0.05) because of the negative correlation in the interval 31700e41000 radiocarbon years BP.

Fig. 4. Comparison of time-related density distributions of mammoth-based 14C dates of “areal” type (New Siberia and Taimyr), and “spot” type: Bykovsky Peninsula, Oyogos-Yar, and Bolshoi Lyakhovsky Island. Indexes after each of the correlation coefficient symbols “r” indicate the pair of samples that have been compared (e.g. “r(L vs. TM)” means the comparison of the Bolshoi Lyakhov Island sample and the Taimyr sample). The first value of the correlation coefficient shows the correlation coefficient of the total curve length, the second value of the correlation coefficient shows the maximal correlation coefficient if we consider different portions of the curves. P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2321

Fig. 5. Comparison of mammoth 14C date frequencies and insect paleoenvironmental proxy record. The black line shows the percentage of xerophilic insect species. The gray line denotes frequencies of 14C mammoth dates from New Siberian Islands and Taimyr Peninsula (combined).

were very productive, do not yield a representative sequence that mammoth in the mainland of the Holarctic.4 Along with many adequately reflects the changing abundance of animals during the other new PHT dates (Table 1) this material suggests a broad interval. Possible reasons include intermittency and irregularity of expansion of the last mainland mammoth refugium to the east, up the terrestrial sedimentation, the low probability of uniform to the 150 E. The details of mammoth extinction will be discussed burial, and exposure rates on a strip of land that is a few kilo- later. meters long. The probability of taphonomic biases decreases when the studied material comes from a wider area and from 3.3. Comparison of the time-distribution of 14C dates on mammoths multiple sites. and climatic proxy data It is evident that a fluctuating trend over a period of tens of thousand years is present in all five distributions. This does not Similar density distribution patterns of mammoth-based contradict the above conclusions. The effect of the largest fluctua- radiocarbon dates from two far-apart regions of the eastern Arctic tions in animal numbers apparently overrides the effect of most demonstrate that the observed fluctuations mostly reflect changes local geological biases. The precise evaluation of the impact of local in animal numbers and are only faintly affected by geological, taphonomic factors on the distribution of dates, however, remains taphonomic, and erosional factors. This conclusion permits us to a difficult task. This is why we regard the results of studies based on search for the causes of fluctuations in numbers of Arctic Siberian all available dates, without applying the quality constraints out- mammoths through the Late Pleistocene and early Holocene. lined above, as debatable (see for example: Kuzmin, 2007, 2010). 3.3.1. Record of changes in insect faunas from the Late Pleistocene deposits of the Lena River delta 3.2. Changes in numbers of mammoths in Arctic Siberia in the Late The comparison of fluctuations in mammoth numbers and the Pleistocene and early Holocene based on time-related density percentage record of xerophilic insects shows a striking covariation 14 distributions of C dates of the curves (Fig. 6). The correlation coefficient of 0.62, p < 0.05, suggests significant dependence of both processes on a third factor. We assume that the clear similarity of the samples originating It is likely that climatic temperature and moisture changes resulted from two Arctic regions (NS and TM), situated more than 1000 km in this effect. fi from each other, provides ample evidence to support the rst These observations also corroborate the broadly accepted “ hypothesis: Under certain sampling conditions the frequency of concept that mammoths strongly depended on a steppe-like radiocarbon dates of mammoth corresponds with the time-related landscape (the xerophilic insect percentage positively depends on ” numbers of this animal. It is evident, that the number of dates can the amount of steppe vegetation). Note that the mammoth record only give an idea about relative, not absolute, animal numbers. terminates at exactly the moment when the percentage of xero- The distribution of dates shows that woolly mammoths popu- philic insects dramatically drops to a very low value (9700 14C ybp). lated vast areas, including the New Siberian Islands and the Taimyr Although there is a short hiatus in the insect record during this region, from some 49 000 until 9470 radiocarbon years BP, or from transition, the insect composition immediately after the event about 55 000 to 10 700 calendar years BP. There were three minima suggests covariation of both factors. of relative mammoth numbers during the last 49 000 years: 22 000, e 14 500 19 000, and 9500 radiocarbon years ago, or around 26 000, 3.3.2. Pollen and insect record of the last 34 ka in the lower Yana e fi 16 20 000, and 10 500 calendar years respectively. Signi cant River, Arctic Siberia e peaks are observed at about 35 000 39 000 radiocarbon years The Yana River record provides fairly detailed and well-dated e e 14 (40 000 45 000 calendar years), and 10 000 14000 C years BP proxy data of absolute temperature and moisture spanning e (11000 15 500 calendar years BP). 9500e34 000 ybp. Although the Yana River site is somewhat to the The collection of youngest radiocarbon dates suggests explosive south of the study area, data suggest that it supplies a rather growth of mammoth numbers both in the New Siberian Islands realistic general picture of past climates across the entire region, (New Siberian Islands were connected to the continent), and in the because during the cold phases of the Pleistocene the latitudinal Taimyr Peninsula, just before the Pleistocene-Holocene transition 14 (PHT). A new C date (GIN-11874) of 9470 40 (10 700 70 4 In Wrangel (Vartanyan et al., 1993, 1995, 2008; Vartanyan, 2007) and Pribilof calendar years BP) indicates the last (known so far) occurrence of Islands (Guthrie, 2004; Veltre et al., 2008) woolly mammoths survived longer. 2322 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Fig. 6. Comparison of mammoth numbers and the Yana River site climatic proxy record. Upper curve, the percentage of xerophilic insect species; middle black curve, the average temperatures of the warmest month; middle gray curve, annual mean precipitation (based on the “environmental envelope” e the environment encompassing the current distribution of a plant taxa (Pitulko et al., 2007, Pavlova et al., 2009)), all parameters presented as deviations from their present-day values. Lower black curve, frequency of mammoth dates from the New Siberian Islands, lower gray curve, frequency of mammoth dates from the Taimyr peninsula. climatic gradient was low. We can infer that general trends in 3.3.4. Lake Baikal core record: concentrations of sponge spicules mammoth numbers are similar to those of the humidity and over time summer temperature (Fig. 6). The comparison of the Lake Baikal climatic proxy record (based The correlation coefficient between 14C numbers and maximal on sponge spicules) and mammoth date frequencies suggests summer temperature is 0.58, p < 0.05. For radiocarbon date a clear agreement between these differently scaled cycles (Fig. 8). numbers and humidity the value is 0.54, p < 0.05. That suggests The correlation coefficient reaches 0.73, p < 0.05. a positive dependence between mammoth numbers and both summer temperature and annual moisture during the colder phase 3.3.5. Greenland ice-core data: GISP2 d18O values of the Late Pleistocene. The smaller-scale fluctuations of GISP2 d18O record do not conform the mammoth date frequencies e the correlation coeffi- 3.3.3. El’gygytgyn Lake paleoenvironments record: change of cient is 0.014 (Fig. 9). However if the GISP2 curve is shifted back- magnetic susceptibility during the last glacial cycle ward by 1100 years (Fig. 9) the conformity in the 13 000e36 000 Comparison of mammoth 14C frequencies and the Lake El’gy- ybp time span is pronounced (r ¼ 0.56, p < 0.05). Atleast four gytgyn (E-Lake) magnetic susceptibility e a proxy for temperature different explanations for this phenomenon can be suggested: and insolation e demonstrates a positive correlation. The correla- tion coefficient is 0.61, p < 0.05 (Fig. 7). Millennial-scale anomalies 1. There is a natural lag between the response of local environ- of the E-Lake record do not fit so closely those of mammoth 14C ments and the global climatic changes. frequencies. This can be interpreted either as a real difference 2. The age model of the GISP2 d18O record is not sufficiently between small-scale climatic changes in the studied regions, or by precise. imprecision in the age model applied to the E-Lake record e two 3. The calibration of the mammoth 14C dates is not correct. age models proposed by Lozkhin et al. (2007) and Nowaczyk et al. 4. There is no dependence between mammoth abundance and (2007) provide scales which differ considerably one from another. climate.

Fig. 7. Comparison between mammoth 14C frequencies and Lake El’gygytgyn magnetic susceptibility, which is believed to be a proxy for temperature and insolation. P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2323

Fig. 8. Comparison of the Lake Baikal climatic proxy record (sponge spicules) and mammoth date frequencies.

We probably can reject the hypotheses 3 and 4, because some of tens of thousands of years-long cycles, are clear in all datasets. the studied climatic proxies fit mammoth 14C frequency distribu- Smaller-scale fluctuations sometimes excellently fit the mammoth tions quite well. Hypotheses 1 and 2 appear to be more credible. In curve, sometimes not. New research is needed to evaluate the any case, cycles tens of thousands of years long (Milankovitch- observed millennial-scale variations in mammoth date distribu- scale) are visible both in the GISP2 d18O record and the Siberian tions. Thus, the comparison of mammoth 14C frequencies (a mammoth date frequencies. reflection of mammoth numbers) and different climatic proxy data from northern Asia, Greenland and the North Atlantic 3.3.6. Mg/Ca ratio in marine ostracod shells from the North Atlantic suggests a positive dependence between mammoth numbers and Comparison of the mammoth date frequency record and the climatic temperature spanning 49 000 to roughly 10 000 calendar Mg/Ca ratio in marine ostracod shells from the North Atlantic years BP. reveals surprisingly high covariation: r ¼ 0.65, p < 0.05 (Fig. 10). This probably reflects a similar response of West Beringian envi- 4. Discussion ronments and the environments in certain parts of North Atlantic, to global climatic changes. The remarkable agreement between the patterns of fluctuation 14 Not all the studied climatic proxies fit mammoth C frequency in mammoth numbers in the Taimyr area and the New Siberian distributions perfectly. This is to be expected as the different Islands (Fig. 3) indicates a common external cause. The strong proxies do not all agree with one another, but least the general, correlation of mammoth numbers with many climatic proxies

Fig. 9. d18O GISP2 Tuned to Hulu (Grootes et al., 1993; Stuiver and Reimer, 1993; Meese et al., 1994a,b) versus mammoth radiocarbon date frequencies. The maximal conformity is reached when GISP2 curve is shifted on 1000 years back position (original position, gray line, shifted, black line). 2324 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Fig. 10. Comparison of the mammoth date frequency record and the Mg/Ca ratio in marine ostracod shells from the North Atlantic. stresses the dominant role of climatic factors. However, the with Milankovitch-scale and also most of the smaller, millennial- response of mammoth populations to the climatic changes was scale temperature changes (Figs. 5e10). Between 10 000 and 9000 not always the same. During the cold and moderately cold phases 14C ybp the trend is reversed e the global increase of temperature of the Late Pleistocene, mammoth numbers correlate positively which caused rise in numbers of mammoth previously, this time,

Fig. 11. Comparison between Arctic Siberia mammoth numbers, North-East Siberia human 14C date frequencies, and the climatic proxy record of Arctic Siberian pollen. Pollen data: Taimyr Peninsula (Andreev et al., 2004); Lena River Delta (Pisaric et al., 2001); New Siberian Islands (Makeyev et al., 2003) demonstrate the most important moment of “slamming shut” of steppe-like landscapes, which occurred 9500 14C ybp all around the studied area. Note that most dated archeological sites occur to the south and to the east of the terminal mammoth population area. P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328 2325 with higher absolute temperatures reached, resulted in a reduc- play an important role as hunted prey. The number of its bones is tion in numbers (Fig. 11). This evidently results from the ecology of much lower than the number of remains of other large herbivores, the mammoth. The insect communities and plant fossil data from particularly horses. the Bykovsky Peninsula (Kienast et al., 2005; Sher et al., 2005), the The archaeological evidence of active human dispersal in lower Yana River (Pitulko et al., 2007; Pavlova et al., 2009), and northern and eastern Siberia becomes clearer with the spread of Taimyr peninsula (Andreev et al., 2003, 2004) provide data on the Dyuktai Culture, roughly correlative to the LGM (Pitulko, 2006). environments and climate spanning up to 55 ka. During the In Northeast Asia, a number of sites e represented by the Dyuktai, moderately cold stage preceding the LGM, arctic plant species Ezhantsy, Verkhnetroitskaya (with reservations), Khaiyrgas, coexisted with steppe, meadow aquatic and littoral taxa. Xero- Avdeikha and, probably, Berelyokh and Ushki I (bed VI) cultures e philic species played an important role in the insect communities. have been radiocarbon dated between 18e20 and 11e12 ka The floristic and insect compositions reflect a continental, semi- (Dikov, 1979; Mochanov, 1977; Mochanov and Fedoseeva, 1975; arid climate with cold low-snow winters, and summers noticeably Kostukevich et al., 1980; Stepanov et al., 2003; Goebel et al., warmer than at present (Guthrie, 1990; Sher, 1997; Yurtsev, 2001). 2003). Notwithstanding relatively few known sites and poor Productive steppe and meadow communities were very favorable knowledge of the vast Arctic region, the existence of a significant for Late Pleistocene herbivores including mammoth. Mammoths human population density by the end of the Pleistocene seems existed in their largest numbers. Further cooling, slightly before probable. People are believed to have populated the valleys of all and especially during the LGM, brought stronger climatic the major rivers in the north and east of Siberia (Pitulko, 2001). Did contrasts, with cool, dry summers and extremely cold winters. the growing density of humans affect the population of These unfavorable conditions decreased the number of animals, mammoths? It is clear that the effect (if any) was quite limited. The but the population quickly recovered when it became even slightly number of dates on mammoths corresponding to the interval from warmer in summer. A relative warming, which started 14 500 15 000 14C ybp to the beginning of the Holocene forms a significant radiocarbon years bp, briefly restored the optimal conditions for peak and then shows an abrupt decline to zero (Fig. 11). The pop- mammoths. However, further very intensive warming sharply ulation of mammoths expands in parallel to the growing number of deteriorated the mammoth’s environment. Plant macrofossil, terminal Paleolithic sites, i.e. increasing number of mammoths with pollen and insect assemblages dated to about 9500 radiocarbon an increasing density of human population. This phenomenon has ybp indicate a sharp paludification possibly caused by atmospheric recently been independently described by Ugan and Bayers (2007). humidity due to marine transgression increasing oceanic influence Thus, the sustainable population of Asian woolly mammoths was (Fig. 11). not observably affected by the presence of humans. This allows us Many steppe-adapted plant and insect taxa vanished from the to reject the “Blitzkrieg overkill” hypothesis of the mammoth area. Productive grass biomes were replaced by low-productivity extinction in Asia (extermination of mammoth by humans shortly taiga and communities. Grazing areas were also shrinking after the first contact). due to greater watering of landscapes, dissection of relief (due to The global restructuring of environments in Arctic Siberia the thermoabrasion of the permafrost ground), and extensive at the PleistoceneeHolocene boundary resulted in shrinkage transgression of the . Furthermore, the increased of the formerly extensive range of mammoth. The youngest winter temperatures and moisture led to higher snow cover. This dates on mammoths have been shown to concentrate in the hampered winter foraging and increased the probability of spring northernmost areas, including Yamal, northern Taimyr and floods, which were sometimes catastrophic (Nikolskiy et al., (Vartanyan et al., 1993, Vartanyan et al., 1995; 2010). This combination of unfavorable conditions caused the Sulerzhitsky, 1997; Stuart et al., 2002). This trend is supported great reversal in the response of mammoth populations to the by new data from the New Siberian Islands and the adjacent climatic change. This time, the increase in annual temperature mainland (this paper). The northern range shift and decrease in reduced mammoth numbers (this paper) and areal size (Stuart body size (see for example Vartanyan et al., 1993; Mol et al., et al., 2002; Orlova et al., 2004). The extreme decrease in the 2006) could have secured survival for the mammoths, but it mammoth populations pushed the species to the brink of did not. It seems plausible that the final extinction of mammoths extinction. occurred due to interactions with humans. Although recent Thus, we consider climatic oscillations as the most important chronological data on the first appearance of humans on the factor in controlling changes in the mammoth population. Climate Taimyr Peninsula, islands of the Arctic (New Siberian Islands, had a direct and indirect influence over the mammoths and drove Wrangel Island), and Pacific (Pribilof Islands) do not strictly them to a critically small population size at the very beginning of support this view e in all cases there is a minor gap between the the Holocene. Were these factors decisive in the ultimate extinction last dates on mammoths and the first dates on humans (Table 2) of Mammuthus primigenius in the Holocene? Extremely low e it is very probable that the contact of “first people” and “last numbers were very probably experienced by the species at various mammoths” occurred in these areas. Both the last mammoths times in the past (Nogués-Bravo et al., 2008). At any rate, large- and the first humans were there so scanty, that their remains are scale warming events, the interglacial periods similar to the Holo- so far paleontologically and archeologically invisible. At least cene, occurred many times in the Pleistocene, yet no extinction data from further south suggest that the human population happened. increased at the same time as the mammoth’s demise further Archeological data need to be taken into consideration. The north, 9500 14Cybp(Fig. 11). appearance of humans in Arctic Siberia at about 28 000 radiocarbon Thus, humans appear not to have been responsible for the years (Pitulko et al., 2004), documented by the Yana Paleolithic Site, destruction of a sustainable mammoth population. The presence evidently did not visibly affect the mammoth (Fig. 3). Although of humans could, however, have been fatal to mammoths during Arctic coastal lowlands were most likely already inhabited, the strong minima in their range and numbers, which occurred at the mammoth population in the north of Siberia would not have very beginning of the Holocene on the mainland and in the middle experienced any serious human pressure because humans were Holocene on Wrangel Island. Humans may have acted as an still very rare in these regions. Moreover, it is not clear if significant additional stress factor, the “last straw”, which destroyed the mammoth mortality occurred due to hunting. The archae- remaining restricted populations of mammoths living in very ozoological record of the Yana Site shows that mammoths did not limited areas. 2326 P.A. Nikolskiy et al. / Quaternary Science Reviews 30 (2011) 2309e2328

Table 2 Comparison of mammoth last occurrence and human first appearance in West Beringia and north Pacific islands.

Area The last occurrence of mammoths (14C BP) The first occurrence of humans (14C BP) Wrangel Island, Bering Sea 3685 60 (Ua-13366), Wrangel Island 3345 70 (Ua-18086), Tchertov Ovrag archeological site, south of (Vartanyan, 2007) Wrangel Island (Gerasimov et al., 2002)a Pribilof Islands, Pacific ocean 5630 40 (Beta 190141B) Late eighteenth century, St. Paul Island (Veltre et al., 2008) St. Paul Island (Veltre et al., 2008) þ polar bear dated to 4430 40 (Beta-182978) New Siberian Islands 9470 40 (GIN-11874), south of 8000e7800, 22 dates, Zhokhov archeological site, New Siberia Island (this report) , (Pitulko, 2003) Taimyr peninsular 9670 60 (GIN-1828), 6030 100 (Le-884), Tagenar VI archeological site, Nizhnyaya Taimyra R. (Stuart, 1991) Central Taimyr (Khlobystin, 1989) Lena-Indigirka coastal lowlands 11 900 50 (GIN-12974), Berelyokh site, 28 250 170 (Beta 173064), Yana archeological site, Lower Indigirka River (Nikolskiy et al., 2010) Lower Yana River (Pitulko et al., 2004) 12 770 170 (GIN-10267), Bykovsky Peninsular, Lena River Delta (Schirrmeister et al., 2002)

a Should be slightly younger due to the basin effect.

5. Conclusions Andreev, A.A., Tarasov, P.E., Siegert, C., Ebel, T., Klimanov, V.A., Melles, M., Bobrov, A.A., Dereviagin, A.Yu., Lubinski, D.J., Hubberten, H.-W., 2003. Late Pleistocene and Holocene vegetation and climate on the northern Taimyr The analysis of a large series of new, reliable radiocarbon dates Peninsula, Arctic Russia. Boreas 32, 484e505. on mammoths from Arctic Siberia, in combination with previously Arslanov, Kh.A., Svezhentsev, Yu.S., 1993. An improved method for radiocarbon e known archives, shows that the frequencies of radiocarbon dates of dating fossil bones. Radiocarbon 35 (3), 387 391. Asikainen, C.A., Francus, P., Brigham-Grette, J., 2007. Sediment fabric, clay miner- randomly-selected mammoth remains can, under certain condi- alogy, and grainsize as indicators of climate change since 65 ka at El’gygytgyn tions, reflect time-related changes in the numbers of these animals. Crater Lake, Northeast Siberia. Journal of Paleolimnology 37 (1), 105e122. 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